Brain control of prehension is thought to rely on two specific brain circuits: a dorsomedial one (involving the areas of the superior parietal lobule and the dorsal premotor cortex) involved in the transport of the hand toward the object and a dorsolateral one (involving the inferior parietal lobule and the ventral premotor cortex) dealing with the preshaping of the hand according to the features of the object. The present study aimed at testing whether a pivotal component of the dorsomedial pathway (area V6A) is involved also in hand preshaping and grip formation to grasp objects of different shapes.Two macaque monkeys were trained to reach and grasp different objects. For each object, animals used a different grip: whole-hand prehension, finger prehension, hook grip, primitive precision grip, and advanced precision grip. Almost half of 235 neurons recorded from V6A displayed selectivity for a grip or a group of grips. Several experimental controls were used to ensure that neural modulation was attributable to grip only.These findings, in concert with previous studies demonstrating that V6A neurons are modulated by reach direction and wrist orientation, that lesion of V6A evokes reaching and grasping deficits, and that dorsal premotor cortex contains both reaching and grasping neurons, indicate that the dorsomedial parieto-frontal circuit may play a central role in all phases of reach-to-grasp action. Our data suggest new directions for the modeling of prehension movements and testable predictions for new brain imaging and neuropsychological experiments.

Reach-to-grasp actions involve several components of forelimb movements needed to direct the hand toward the object to be grasped, and to orient and preshape the hand according to the object axis and shape. Area V6A, which represents a node of the dorsomedial frontoparietal circuits, has so far been implicated only in directing the arm toward different spatial locations. The present results confirm this finding and demonstrate, for the first time, that during reach-to-grasp, V6A neurons are also modulated by the orientation of the hand. In the present work the object to be grasped was a handle that could have different orientations. Reach-to-grasp movements were executed in complete darkness while gazing at a small fixation point. The majority of the tested cells (76/142; 54%) turned out to be sensitive to the orientation of the handle. Neurons could be modulated during preparation or execution of reach-to-grasp movements. The most represented cells were those modulated by hand orientation both during preparatory and movement periods. These data show that reaching and grasping are processed by the same population of neurons, providing evidence that the coordination of reaching and grasping takes place much earlier than previously thought, i.e., in the parieto-occipital cortex. The data here reported are in agreement with results of lesions to the medial posterior parietal cortex in both monkeys and humans, and with recent imaging data in humans, all of them indicating a functional coupling in the control of reaching and grasping by the medial parietofrontal circuit.

The visuomotor medial posterior parietal area V6A has been recently subdivided into two cytoarchitectonic sectors called V6Ad and V6Av (Luppino et al., 2005). The aim of the present study was to recognize whether these two cortical sectors show different functional profiles. Fourteen hemispheres from eight animals (Macaca fascicularis) were included in this study, for a total of 3828 extracellularly recorded neurons assigned to areas V6Ad or V6Av on cytoarchitectural basis. The sensitivity of recorded neurons to sensory- and motor-related activities was checked with a series of functional tests performed on behaving animals. We found that cells sensitive to visual stimuli were more represented in V6Av and cells sensitive to somatosensory stimuli were more represented in V6Ad. Visual cells directly encoding spatial locations (real-position cells) were present only in V6Av. Cells encoding basic visual and somatic properties as well as different aspects of reaching and grasping activities were present in both sectors of V6A, although with different incidence. Cells that had reach-related activity enhanced by visual feedback and grasping neurons activated by whole-hand prehension were more concentrated in V6Av. Conversely, reaching neurons inhibited by visual feedback and grasping neurons activated by precision grip were more represented in V6Ad. Although V6Av and V6Ad show partly different functional profiles, our data support the idea that V6A is a single functional area involved in the control of reach-to-grasp movements, with the dorsal sector (V6Ad) more involved in the somatomotor control and the ventral sector (V6Av) in the visual control of reaching and grasping actions.

Parietal area V6A contains neurons modulated by the direction of gaze as well as neurons able to code the direction of arm movement. The present study was aimed to disentangle the gaze effect from the effect of reaching activity upon single V6A neurons. To this purpose, we used a visuomotor task in which the direction of arm movement remained constant while the animal changed the direction of gaze. Gaze direction modulated reach‐related activity in about two‐thirds of tested neurons. In several cases, modulations were not due to the eye‐position signal per se, the apparent eye‐position modulation being just an epiphenomenon. The real modulating factor was the location of reaching target with respect to the point gazed by the animal, that is, the retinotopic coordinates towards which the action of reaching occurred. Comparison of neural discharge of the same cell during execution of foveated and non‐foveated reaching movements, performed towards the same or different spatial locations, confirmed that in a part of V6A neurons reaching activity is coded retinocentrically. In other neurons, reaching activity is coded spatially, depending on the direction of reaching movement regardless of where the animal was looking at. The majority of V6A reaching neurons use a system that encompasses both of these reference frames. These results are in line with the view of a progressive visuomotor transformation in the dorsal visual stream, that changes the frame of reference from the retinocentric one, typically used by the visual system, to the arm‐centred one, typically used by the motor system.